Testudo graeca

Superregnum: Eukaryota
Regnum: Animalia
Subregnum: Eumetazoa
Cladus: Bilateria
Cladus: Nephrozoa
Superphylum: Deuterostomia
Phylum: Chordata
Cladus: Craniata
Subphylum: Vertebrata
Infraphylum: Gnathostomata
Superclassis: Tetrapoda
Cladus: Reptiliomorpha
Cladus: Amniota
Classis: Reptilia
Cladus: Eureptilia
Cladus: Romeriida
Subclassis: Diapsida
Cladus: Sauria
Cladus: Archelosauria
Division: Pan-Testudines
Division: Testudinata
Ordo: Testudines
Subordo: Cryptodira
Superfamilia: Testudinoidea

Familia: Testudinidae
Genus: Testudo
Subgenus: Testudo
Species: Testudo graeca
Subspecies: T. g. anamurensis – T. g. antakyensis – T. g. armenica – T. g. buxtoni – T. g. flavominimaralis – T. g. floweri – T. g. graeca – T. g. ibera – T. g. lamberti – T. g. marokkensis – T. g. nabeulensis – T. g. nikolskii – T. g. pallasi – T. g. soussensis – T. g. terrestris – T. g. zarudnyi
Name

Testudo graeca Linnaeus, 1758

Synonyms

Testudo pusilla Linnaeus, 1758

References
Primary references

Linnaeus, C. 1758. Systema Naturae per regna tria naturæ, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis, Tomus I. Editio decima, reformata. Holmiæ: impensis direct. Laurentii Salvii. i–ii, 1–824 pp DOI: 10.5962/bhl.title.542: 198. Reference page.

Additional references

Ananjeva, N.B., Milto, K.D., Barabanov, A.V. & Golynsky, E.A. 2010. An annotated type catalogue of amphibians and reptiles collected by Nikolay A. Zarudny in Iran and Middle Asia. Zootaxa 4722(2): 101–128. DOI: 10.11646/zootaxa.4722.2.1. Paywall. Reference page.

Vernacular names
azərbaycanca: Aralıq dənizi tısbağası
български: Шипобедрена костенурка
català: Tortuga grega
čeština: Želva žlutohnědá
dansk: Maurisk landskildpadde
Deutsch: Maurische Landschildkröte
English: Spur-thighed Tortoise, Greek Tortoise
español: Tortuga mora
فارسی: لاک‌پشت مهمیزدار
suomi: Espanjankilpikonna
français: Tortue grecque
galego: Tartaruga moura
עברית: צב יבשה מצוי
hrvatski: Grčka čančara
magyar: Mór teknős
հայերեն: Կրիա միջերկրածովային
italiano: Tartaruga greca, tartaruga moresca
日本語: ギリシャリクガメ
lietuvių: Graikinis vėžlys
македонски: Полска желка
Nederlands: Moorse landschildpad
polski: Żółw śródziemnomorski, Żółw iberyjski
پنجابی: یونانی کھچوپڑا
português: Testudo graeca
русский: Средиземноморская черепаха
slovenčina: Korytnačka žltohnedá
svenska: Morisk landsköldpadda
Türkçe: Mahmuzlu Akdeniz kaplumbağası
українська: Середземноморська черепаха

The Greek tortoise (Testudo graeca), also known commonly as the spur-thighed tortoise,[1] is a species of tortoise in the family Testudinidae. Testudo graeca is one of five species of Mediterranean tortoises (genera Testudo and Agrionemys). The other four species are Hermann's tortoise (T. hermanni), the Egyptian tortoise (T. kleinmanni), the marginated tortoise (T. marginata), and the Russian tortoise (A. horsfieldii). The Greek tortoise is a very long-lived animal, achieving a lifespan upwards of 125 years, with some unverified reports up to 200 years.[3]

Geographic range

The Greek tortoise's geographic range includes North Africa, Southern Europe, and Southwest Asia. It is prevalent in the Black Sea coast of the Caucasus (from Anapa, Russia, to Sukhumi, Abkhazia, Georgia, to the south), as well as in other regions of Georgia, Armenia, Iran, and Azerbaijan.
Evolution

The oldest known definitive fossil is from the Early Pliocene of Greece,[4] but specimens referred to as Testudo cf. graeca are known from the Late and Middle Miocene in Greece and Turkey.[5][6]
Characteristics

The Greek tortoise (T. g. ibera) is often confused with Hermann's tortoise (T. hermanni ). However, notable differences enable them to be distinguished.
T. g. ibera
T. graeca, male

Greek tortoise	Hermann's tortoise
Large symmetrical markings on the top of the head	Only small scales on the head
Large scales on the front legs	Small scales on the front legs
Undivided carapace over the tail	Tail carapace almost always divided
Notable spurs on each thigh	No spurs
Isolated flecks on the spine and rib plates	Isolated flecks only on the spinal plates
Dark central fleck on the underside	Two black bands on the underside
Shell somewhat oblong rectangular	Oval shell shape
Widely stretched spinal plates	Small spinal plates
Movable posterior plates on underside	Fixed plates on underside
No tail spur	Tail bears a spur at the tip

Subspecies
Testudo graeca, 4 years

The division of the Greek tortoise into subspecies is difficult and confusing. Given their huge range over three continents, the various terrains, climates, and biotopes have produced a huge number of varieties, with new subspecies constantly being discovered. Currently, at least 20 subspecies are published:

T. g. graeca (North Africa and South Spain)
T. g. soussensis (South Morocco)
T. g. marokkensis (North Morocco)
T. g. nabeulensis - Tunisian tortoise (Tunisia)
T. g. cyrenaica (Libya)
T. g. ibera (Turkey)
T. g. armeniaca - Armenian tortoise (Armenia)
T. g. buxtoni (Caspian Sea)
T. g. terrestris (Israel/Lebanon)
T. g. zarudnyi (Iran/Azerbaijan)
T. g. whitei (Algeria)
T. g. floweri (Jordan)

This incomplete listing shows the problems in the division of the species into subspecies. The differences in form are primarily in size and weight, as well as coloration, which ranges from dark brown to bright yellow, and the types of flecks, ranging from solid colors to many spots. Also, the bending-up of the edges of their carapaces ranges from minimal to pronounced. So as not to become lost in the number of subspecies, recently, a few tortoises previously classified as T. graeca have been assigned to different species, or even different genera.

The genetic richness of T. graeca is also shown in its crossbreeding. Tortoises of different form groups often mate, producing offspring with widely differing shapes and color. Perhaps the best means of identification for the future is simply the place of origin.

The smallest, and perhaps the prettiest, of the subspecies, is the Tunisian tortoise. It has a particularly bright and striking coloration. However, these are also the most sensitive tortoises of the species, so they cannot be kept outdoors in temperate climates, as cold and rainy summers quickly cause the animals to become ill. They are also incapable of long hibernation.

At the other extreme, animals from northeastern Turkey are very robust, such as Hermann's tortoise. The largest specimens come from Bulgaria. Specimens of 7 kg (15 lb) have been reported. In comparison, the Tunisian tortoise has a maximum weight of 0.7 kg (1.5 lb). T. graeca is also closely related to the marginated tortoise (T. marginata). The two species can interbreed, producing offspring capable of reproduction.
Sexing

Males of T. graeca differ from females in six main points. Firstly, they are generally smaller. Their tails are longer than females and taper to a point evenly, and the cloacal opening is farther from the base of the tail. The underside is somewhat curved, while females have a flat shell on the underside. The rear portion of a male's carapace is wider than it is long. Finally, the posterior plates of the carapace often flange outward.
Mating and reproduction
A pair of Testudo graeca mating in Mountain Yamanlar Nature Park, İzmir Province, Turkey

In T. graeca, immediately after waking from hibernation, the mating instinct starts up. The males follow the females with great interest, encircling them, biting them in the limbs, ramming them, and trying to mount them. During copulation, the male opens his mouth, showing his red tongue and making squeaking sounds.

During mating, the female stands still, bracing herself with her front legs, moving the front part of her body to the left and right in the same rhythm as the male's cries. One successful mating will allow the female to lay eggs multiple times. When breeding in captivity, the pairs of females and males must be kept separate. If multiple males are in a pen, one takes on a dominant role and will try to mate with the other males in the pen. If more males than females are in a pen, the males might kill each other to mate with the females.

One or two weeks before egg laying, the animals become notably agitated, moving around to smell and dig in the soil, even tasting it, before choosing the ideal spot to lay the eggs. One or two days before egg laying, the female takes on an aggressive, dominant behavior, mounting another animal as for copulation and making the same squeaking sound the male produces during copulation. The purpose of this behavior is to produce respect in the tortoise community so that the female will not be disturbed by the others during egg laying. Further details of egg-laying behavior are the same as those detailed for the marginated tortoise.
Trade

The Greek tortoise is commonly traded as a pet in source countries such as Morocco and Spain, despite the illegality of this trade.[7][8][9] This can lead to an unsustainable removal of wild individuals for the local pet trade and for export. Also, welfare concerns exist with this trade, as the animals are not properly housed when being sold, causing a high rate of mortality in captivity.[10]
Food

The Greek tortoise loves dandelion leaves and other leafy plants. However, although they also enjoy eating lettuce, it is not recommended to them due to having a lack of nutrients that the tortoises need to survive.[11]
See also

List of reptiles of Italy
Mediterranean tortoise
Timothy (tortoise)
Jackson ratio

References

Tortoise.; Freshwater Turtle Specialist Group (1996). "Testudo graeca". IUCN Red List of Threatened Species. 1996: e.T21646A9305693. doi:10.2305/IUCN.UK.1996.RLTS.T21646A9305693.en. Retrieved 19 November 2021.
Fritz, Uwe; Havaš, Peter (2007). "Checklist of Chelonians of the World" (PDF). Vertebrate Zoology. 57 (2): 296–300. ISSN 1864-5755. Archived (PDF) from the original on 17 December 2010. Retrieved 29 May 2012.
"Testudo graeca". The Moirai – Aging Research. 12 September 2016. Retrieved 13 December 2016.
Vlachos E (2015). "The Fossil Chelonians of Greece. Systematics - Evolution - Stratigraphy - Palaeoecology". Scientific Annals of the School of Geology, Aristotle University of Thessaloniki, Greece. 173: 1–479.
Vlachos E., Tsoukala E. (2014). "Testudo cf. graeca from the new Late Miocene locality of Platania (Drama basin, N. Greece) and a reappraisal of previously published specimens". Bulletin of the Geological Society of Greece. 48: 27–40.
Staesche K., Karl H.V., Staesche U. (2007). "Fossile Schildkroten aus der Turkei. In U. Staesche (ed.)". Fossile Schildkroten aus drei Kontinenten. 98: 91–149.
Pérez, I.; Tenza, A.; Anadón, J.D.; Martínez-Fernández, J.; Pedreño, A.; Giménez, A. (2012). "Exurban sprawl increases the extinction probability of a threatened tortoise due to pet collections". Ecological Modelling. 245: 19–30. doi:10.1016/j.ecolmodel.2012.03.016. hdl:10261/67281.
Bergin & Nijman (2014). "Open, Unregulated Trade in Wildlife in Morocco's Markets, TRAFFIC Bulletin". Retrieved 23 March 2015.
Nijman, V; Bergin, D (2017). "Trade in spur-Thighed tortoises Testudo graeca in Morocco: Volumes, value and variation between markets". Amphibia-Reptilia.
Bergin, D.; Nijman, V (2018). "An Assessment of Welfare Conditions in Wildlife Markets across Morocco". Journal of Applied Animal Welfare Science.
"Helpfull [sic] advice for your tortoise diet". www.tortoisecentre.co.uk. Retrieved 29 January 2018.